taxonID	type	description	language	source
039B87A2FFFB3B53D7F2FE07FC96FD05.taxon	type_taxon	Type species: Trox horridus Fabricius, 1775: 818 (by monotypy).	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFFB3B51D7F2FCDEFD36FDD6.taxon	description	(Figs 1 – 5, 18, 21, 24)	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFFB3B51D7F2FCDEFD36FDD6.taxon	materials_examined	Type locality. South Africa, Western Cape province, Klaarstroom 18 km N, Zwartskraal Farm, 33 ° 10 ′ S 22 ° 32 ′ E; verbatim as in neotype label data below. This place lies in the very south of the Great Karoo, at an estimated elevation of 890 – 920 m above mean sea level. Type material examined. Neotype ♂, here designated (TMSA, aedeagus extracted) (Figs 1 – 5): Rectangle, white, with printed text: “ S. Afr., Cape-Karoo | Zwartskraal farm | 33.10 ′ S- 22. ° 32 ′ E [33 ° 10 ′ S 22 ° 32 ′ E] ” || Rectangle, white, with printed text: “ 1.2.1979; E-Y: 1545 | groundtraps [pitfall traps], 29 days | leg. R. Oostrhuizen [recte Oosthuizen] ” || Rectangle, white, with printed text: “ groundtraps with | meat bait ” || Rectangle, white, with printed text: “ TROX | sulcatus | Thunberg | det C. H. Scholtz ” || Rectangle, white, with printed text: “ Phoberus sulcatus | (Thunberg, 1787) | det. WP Strümpher, 2022 ” || Rectangle, red, with printed text: “ NEOTYPE: Phoberus | sulcatus (Thunberg, 1787) | des. W. P. Strümpher, 2025 ” || Small rectangle, white, with printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000934 ”.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF93B5CD7F2FD0AFE55F8FA.taxon	description	The catalogue of insect type specimens in the Thunberg collection at Uppsala University (UUZM) (Wallin & Wallin 2001) lists two “ type ” specimens (Nos 2140 and 2160) of Throx sulcatus. Upon examination of the corresponding material, it was discovered that the two specimens in the collection presently listed as types of Throx sulcatus are, in fact, specimens of Phoberus horridus (Fabricius, 1775) (Fig. 25). The latter is a large, easily recognisable, flightless species found in the southwestern part of the Western Cape province of South Africa (Scholtz, 1980). Johan Christian Fabricius described it (originally in the genus Trox) 12 years before Thunberg described Throx sulcatus. Thunberg described Trox silphoides (also as Throx) on the same 1787 page. Harold (1872: 36) synonymized the latter nominal species with Phoberus horridus (as Trox horridus), an action with which we agree; this is the species masquerading as Phoberus sulcatus in the Thunberg collection (Fig. 25). In his more detailed redescription of Trox silphoides, Thunberg (1818: 450) mentioned that the body is dorsally convex (“ supra convexus ”) and the elytra are closed (“ Elitra clause ”) (here interpreted as being fused) — the main characteristics typically reflected in the external morphology of flightless trogid species, such as Phoberus horridus, are tightly fused elytra and a body shape convex in lateral view. Phoberus sulcatus, on the other hand, has fully functional wings, has a flatter general body profile, and its elytra separate easily; Thunberg (1818: 449) explicitly stated that this species is winged (“ alato ”). It is notable that Throx sulcatus and Throx silphoides were both described on page 38 of Thunberg (1787), and are the only two nominal species of Phoberus (or Trox) described in that work. We are certain that Thunberg would not have confused Trox sulcatus with Trox silphoides, and that the two specimens of Phoberus horridus standing as such (Fig. 25) cannot be (the) syntypes of Trox sulcatus. Both Phoberus sulcatus and Phoberus horridus are morphologically well defined, readily recognisable as different species. We suspect that the type specimens of Trox sulcatus were mistakenly substituted with specimens of Phoberus horridus (i. e. Trox silphoides) in the Thunberg collection. It is clear that the red curatorial type labels (Fig. 25) currently associated with the two Phoberus horridus specimens were added more recently, as they were produced by digital printing. Labels pinned in the drawers that contained the specimens, handwritten by Thunberg (handwriting compared in Horn et al. 1990: Teil II, plate 36, fig 17), are the only remaining evidence of the Throx sulcatus types having been in Thunberg’s collection (Fig. 25). There are no other specimens of Trox sulcatus present in the Thunberg collection. Our efforts to locate the missing syntypes in other museums (MNHB, MNHN, NHMB, NHMUK, NHMW, NHRS, NMPC, SDEI) were unsuccessful. We hence believe that the original type material of Throx sulcatus is lost. An equivalent case of mistaken substitution was also noted for one of the two syntypes of Throx silphoides (Thunberg type No. 2142) in the Thunberg collection, which appears to have been substituted with a specimen of Phoberus caffer (Harold, 1872) (WPS, personal observation). The Thunberg collection in Uppsala contains inconsistencies due to mistaken substitutions that require further investigation. Such substitutions are particularly likely in this collection because none of the specimens have locality labels, determination labels or labels indicating their type status. Additionally, all the handwritten name labels are pinned inside the drawers or trays instead of being attached to the specimens. This practice was not uncommon in historical collections from that era. A similar situation was recently highlighted by Costa-Silva et al. (2024) in the Joseph Banks collection in NHMUK. Even though the types of Trox sulcatus could not be studied / traced, Harold (1872), Haaf (1953) and Scholtz (1979, 1980) clearly defined the distinguishing characteristics of Phoberus sulcatus (then as Trox), which can be readily identified from other similar species in the genus by the characteristic male genitalia (Figs 4, 18, 21). Type specimens are important to maintain the nominal species’ stability and to avoid future misidentification. Considering that the supposed missing type specimens of Trox sulcatus from Thunberg’s collection correspond to the modern concept of Phoberus sulcatus (sensu Harold [1872], Haaf [1953] and Scholtz [1979,1980]), we understand that a designation of a neotype is necessary according to Article 75 of the ICZN (1999). Therefore, we propose the designation of a neotype here to establish the single name-bearing type specimen of Throx sulcatus. The neotype specimen fulfils the requirements of ICZN Article 75.3 (ICZN 1999). We selected a well-preserved male from Zwartskraal farm in the Western Cape province, South Africa, from the collection of the TMSA (Figs 1 – 5).	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF93B5CD7F2FD0AFE55F8FA.taxon	materials_examined	Additional material with precise localities examined (599 specimens). SOUTH AFRICA: Western Cape. 1 ♂ [Matjiesfontein], Table Farm, 33 ° 10 ′ S 20 ° 31 ′ E, found on carcass, 2. iii. 1996, KI Krel (CJBH). 2 ♂, 2 ♀ Beaufort West, De Hoop [farm], 32 ° 08 ′ S 22 ° 44 ′ E, 14 – 16. ix. 1983, S Louw & A van Rensburg (BMSA). 11 ♂, 13 ♀ Klaarstroom, Zwartskraal [farm], 33 ° 10 ′ S 22 ° 32 ′ E, riverine bush, pitfall traps: 8. xi. 1978 - 16. i. 1979, E-Y: 1540, S Endrödy-Younga (TMSA). 3 ♂, 28 spec. unsexed idem but 31. iii. 1979 - 29. iv. 1979, E-Y: 1568 (TMSA). 1 ♀ idem but 30. viii. 1979, E-Y: 1622, hand-collected, sand (TMSA). 18 ♂, 116 spec. idem but 1. ii. 1979 - 1. iii. 1979, E-Y: 1545, R Oosthuizen (1 SAMC; 133 TMSA). 12 ♂, 18 spec. unsexed idem but 2. iii. 1979 - 30. iii. 1979, E-Y: 1552 (TMSA). 2 ♂, 9 spec. unsexed idem but 30. iv. 1979 - 10. vi. 1979, E-Y: 1571 (TMSA). 1 ♂, 23 spec. unsexed idem but 11. vi. 1979 - 22. vii. 1979, E-Y: 1574 (TMSA). 4 ♂, 5 ♀ idem but rocky ridge, 23. vii. 1979 - 4. ix. 1979, E-Y: 1637 (TMSA). 2 ♂, 4 ♀ idem but 5. ix. 1979 - 24. x. 1979, E-Y: 1639 (TMSA). 4 ♂, 2 ♀ idem but 25. x. 1979 - 8. xii. 1979, E-Y: 1672 (TMSA). 4 ♂, 2 ♀ idem but 15. xii. 1979 - 29. i. 1980, E-Y: 1693 (TMSA). 12 ♂, 18 spec. unsexed idem but E-Y: 1699 (TMSA). 8 spec. unsexed idem but 18. iii. 1980 - 5. v. 1980, E-Y: 1702 (TMSA). 2 ♂, 5 ♀ idem but E-Y: 1707 (TMSA). 8 spec. unsexed idem but 26. vi. 1980 - 5. viii. 1980, E-Y: 1710 (TMSA). 1 ♀ idem but hand-collected at farm house, 26. vi. 1980, E-Y: 1715 (TMSA). 23 spec. unsexed idem but 22. ix. 1980 - 7. xi. 1980, E-Y: 1717 (TMSA). 15 spec. unsexed idem but 8. xi. 1980 - 19. xii. 1980, E-Y: 1737 (TMSA). 1 ♂, 1 ♀ Zwartskraal farm, 33 ° 10 ′ S 22 ° 32 ′ E, x. 1983, R Oosthuizen (TMSA). 1 ♂ Groot Swartberg Nature Reserve, 33 ° 20 ′ S 22 ° 01 ′ E, i – ii. 2006, Z van der Walt (SANC). 4 ♂, 4 ♀ Oudtshoorn, Swartberg Pass, 33 ° 21 ′ S 22 ° 03 ′ E, pitfall traps: 13. iii. 1979, W Breytenbach (TMSA). 5 ♂, 4 ♀ Oudtshoorn, Koos Raubenheimer Dam, 2 km E, 33 ° 24 ′ S 22 ° 19 ′ E, bottom of dry, stony hill, pitfall traps: 22. x. 1993 - 5. xi. 1993, E-Y: 2892, S Endrödy-Younga (TMSA). Swartberg, Meiringspoort [ca. 33 ° 25 ′ S 22 ° 33 ′ E], ii. 1932, KH Barnard (SAMC). 1 ♂ Swartberg, Meiringspoort, Spitskop, [ca. 33 ° 28 ′ S 22 ° 33 ′ E], i. 1935, Museum staff (NHMB). 1 ♂ 2 spec. unsexed De Vlug, near Avontuur, Prince Alfred Pass, [ca. 33 ° 45 ′ S 23 ° 08 ′ E], 800 m, 16 – 19. xi. 1981, fynbos, S & J Peck (CMNC). 1 ♂ Wilderness, [ca. 33 ° 59 ′ 40 ″ S 22 ° 34 ′ 30 ″ E], xii. 1955, V Fitzsimons (TMSA). Northern Cape. 1 ♂ Victoria West, Melton Wold 158 [farm, mine], [31 ° 27 ′ S 22 ° 45 ′ E], 8 – 10. iii. 1989, Entomology Dept. (BMSA). 2 ♀ Richmond, Mynfontein [farm], [ca. 30 ° 55 ′ S 23 ° 56 ′ E], vi. 1971, JS van Zyl (TMSA). Eastern Cape. 2 ♀ Joubertina [33 ° 49 ′ S 23 ° 51 ′ E], 17. i. 1982, AE Oelofsen (BMSA). 1 ♂ Tsitsikamma National Park, 34 ° 01 ′ S 23 ° 53 ′ E, iii. 1992, J Klimaszewski (TMSA). 1 ♂ Aberdeen [32 ° 29 ′ S 24 ° 04 ′ E], i. 1953, J Heyns (SAMC). 1 ♂, 2 ♀ Graaff-Reinet, Paardekraal [farm], 32 ° 12 ′ S 24 ° 09 ′ E, 1,002 m, 3 – 7. ii. 2003, dry xeric slope, DM Kroon (TMSA). 1 spec. unsexed Baviaanskloofberg, 33 ° 36 ′ S 24 ° 23 ′ E, 868 m, pitfall traps: 15. xi. 1982 - 15. xii. 1982, WB 238, W Breytenbach (TMSA). 5 ♀ idem but 595 m, WB 240 (TMSA). 2 ♂, 2 ♀ idem but 183 m, WB 241 (TMSA). 1 ♂ Baviaanskloof Nature Reserve, 33 ° 38 ′ S 24 ° 14 ′ E, pitfall traps: 24. i. 2021, WP Strümpher & GM Daniel (TMSA). 1 ♀ Sneeuberge, Petersburg, 32 ° 19 ′ S 24 ° 58 ′ E, 6 – 14. xi. 2012, M Krüger (TMSA). 8 unsexed Middelburg, 31 ° 30 ′ S 25 ° 01 ′ E, pitfall traps: viii – x. 1984, R Loubser (SANC). 2 ♂, 8 ♀ idem but x – xii. 1985, M de Jager (SANC). 6 ♂, 6 ♀ idem but i – xii. 1986 (SANC). 4 ♂, 5 ♀ Nxuba, Mountain Zebra National Park, [ca. 32 ° 15 ′ S 25 ° 27 ′ E], 30. x. 1986, A van Rensburg (BMSA). 1 spec. unsexed Hogsback, 33 ° 35 ′ S, 26 ° 55 ′ E, collected on light trap, 16. x. 2004, SI Wilsher (CJBH). 4 ♂, 1 ♀ Gqeberha, [ca. 33 ° 57 ′ S 25 ° 36 ′ E], Dr Brauns, [no date] (TMSA). 1 ♂ Gqeberha, Uitenhage Farms, Witteklip [33 ° 54 ′ S 25 ° 15 ′ E], JL Drege (SANC). 1 ♂ Suurberg Pass, [ca. 33 ° 17 ′ S 25 ° 43 ′ E], 21. ii. 1954, L. Vári (TMSA). 1 ♀ Addo Elephant National Park, [ca. 33 ° 28 ′ S 25 ° 47 ′ E], 4. x. 1996, KS Coles (TMSA). 1 spec. unsexed Addo Elephant National Park, Kabouga House, 33 ° 18 ′ S 25 ° 22 ′ E, 217 m, x. 2019 (CJBH). 11 ♂, 4 ♀ Middleton, Sheldon [station], [33 ° 01 ′ S 25 ° 50 ′ E], 8. i. 1950, Zumpt (NHMB). 1 ♂, 1 ♀ Tarkastad, Kommandodrif Nature Reserve, 32 ° 07 ′ S 26 ° 02 ′ E, pitfall traps: 1 – 3. ii. 2021, WP Strümpher & GM Daniel (TMSA). 10 ♂, 10 ♀, 31 unsexed Sidbury, Amakhala Game Reserve, 33 ° 31 ′ 16 ″ S 26 ° 08 ′ 27 ″ E, pitfall traps baited with carnivore dung: iii – iv. 2016, C Tocco (AMGS). 1 ♀ Makhanda SW, Coldsprings [lodge], [33 ° 20 ′ S 26 ° 29 ′ E], 17. ii. 1976, CF Jacot-Guillarmod (AMGS). 1 ♂ Thomas Baines Nature Reserve, 33 ° 24 ′ S 26 ° 30 ′ E, 5. v. 2005, JM Midgley, in exposed small intestine of dead White Rhinoceros Ceratotherium simum (Burchell) (TMSA). 1 spec. unsexed Thomas Baines Nature Reserve, 33 ° 25 ′ S 26 ° 30 ′ E, 25. iii. 1996, C Mcilwaine (CJBH). 2 ♂, 1 ♀ Makhanda, 33 ° 20 ′ S 26 ° 32 ′ E, 648 m, 5. ix. 2009, WP Strümpher (TMSA). 3 ♀ Makhanda, 1893 (1 SAMC, 2 SANC). 2 ♂, 2 ♀ Makhanda, May [18] 97, Dr Chew (3 AMGS; 1 SAMC). 1 ♂ Makhanda, 6. viii. 1967, M Bruton (TMSA). 1 spec. unsexed Makhanda, 33 ° 18 ′ S 26 ° 31 ′ E, 10. v. 2003, PD Collett (CJBH). 1 spec. unsexed Makhanda, 33 ° 19 ′ S, 26 ° 32 ′ E, 12. v. 2005, L De Wet (CJBH). 1 ♀ Makhanda, 33 ° 19 ′ S 26 ° 32 ′ E, 20. iv. 2005, JM Midgley, on dead Black-backed Jackal Lupulella mesomelas (Schreber) (TMSA). 2 ♂, 2 ♀ Makhanda, Hilton, [33 ° 15 ′ S 26 ° 21 ′ E], 11. x. 1983, AJ Weaving, from dried carcass of Vervet Monkey (AMGS). 1 ♂, 1 ♀ idem but 16. x. 1983 (AMGS). 1 ♀ idem but 4. xii. 1983 (AMGS). 1 ♂ Makhanda, Rhodes University Campus, 33 ° 18 ′ S 26 ° 32 ′ E, 17. vi. 1976, RHG Clarke (TMSA). 5 ♂, 5 ♀, 20 unsexed Makhanda, Signal Hill, 33 ° 20 ′ S 26 ° 33 ′ E, 17. ix. 2014, JM Midgley & MH Villet, on dead jackal, 1 week (AMGS). 2 ♂, 2 ♀, 8 unsexed idem but 8. x. 2014, on dead jackal, 4 weeks (AMGS). 3 ♂, 3 ♀, 14 unsexed idem but 20. x. 2014, on dead jackal, 6 weeks (AMGS). 2 ♂ Katberg [32 ° 29 ′ S 26 ° 41 ′ E], 1 – 13. xi. 1932 (TMSA). 1 ♀ idem but 4 000 ft. [ca. 1,220 m], xii. 1932 (TMSA). 1 ♂ idem but 19 – 26. ii. 1933 (TMSA). 1 ♂ Komani, [31 ° 54 ′ S 26 ° 52 ′ E], i. 1975, R. W. (SAMC). 1 ♀ Port Alfred, Kap River Nature Reserve, 33 ° 29 ′ S 27 ° 07 ′ E, 4. ix. 1994, on dried remains of Cape Bushbuck Tragelaphus sylvaticus (Sparrman) carcass, AJ Gardiner & EJ Bruce-Miller (TMSA). 1 ♂ NW of Stutterheim, 32 ° 32.3 ′ S 27 ° 23.5 ′ E, 17 – 20. i. 2016, P Bulirsch (CJBH). 1 ♂ East London, 33 ° 01 ′ S 27 ° 55 ′ E, x. 1974, GJ Petty (SANC). 1 ♂ East London, Suffolk Farm, 5. xii. 1998, PG Barratt (CJBH). 1 ♂ Mthatha, 31 ° 35 ′ S 28 ° 47 ′ E, 4. xii. 1989, N Duke (SANC). KwaZulu-Natal. 1 ♀ Durban, [ca. 29 ° 51 ′ S 31 ° 01 ′ E], 1894, N Barker (SAMC). 1 ♀ Umlaas Road, 29 ° 44 ′ S 30 ° 30 ′ E, 866 m, i. 1983, P Atkinson (SAMC). 1 ♂ idem but iv. 1983 (SAMC). 1 ♂, 1 ♀ idem but x. 1984 (SANC). 1 ♀ idem but xii. 1984 (SANC). 1 ♀ idem but x. 1985 (SAMC). 1 unsexed idem but vi. 1997, PE Reavell (SANC). 1 ♂, 1 ♀ Pietermaritzburg, [ca. 29 ° 36 ′ S 30 ° 22 ′ E], x. 1983, PE Reavell (SAMC). 1 ♂ Wartburg, [29 ° 26 ′ S 30 ° 34 ′ E], 11. ii. 1953, IH Wiese (SANC). 1 ♂, 1 ♀ oThongathi NNW, Warwick farm, 29 ° 25 ′ S 31 ° 04 ′ E, vi. 2002, PE Reavell (SANC). 1 ♀ Dalton, 29 ° 20 ′ S 30 ° 37 ′ E, vi. 1997, PE Reavell (SANC). 2 ♂, 3 ♀ idem but xii. 1997 (TMSA). 1 ♂ Estcourt, [29 ° 01 ′ S 29 ° 52 ′ E], Haviland (SAMC). 1 unsexed Richards Bay, Lake Chubhu, 28 ° 52 ′ S 31 ° 58 ′ E, 19. iii. 1980, light trap, RG Oberprieler (SANC). Additional material without precise locality examined (34 specimens). SOUTH AFRICA. 2 ♂, 3 unsexed Cap. b. Esp. [Cap de Bonne-Espérance, Cape of Good Hope] (MNHN). 1 ♂ Cape Colony, 06. vi. 1881 (SAMC). 2 unsexed Cap [Cape Colony], Schmidt, Coll H Schapler (SDEI: # 303411, # 303413). 1 unsexed Cap, Péringuey (MNHN). 3 ♂, 2 ♀, 3 unsexed Cap (MNHN). 1 ♂ Le Cap (MNHN). 2 ♂ Cape (MNHN). 1 unsexed Caffraria, Coll Kraatz (SDEI: # 303412). 1 ♂, 1 unsexed South Africa (SAMC). 1 unsexed Afrique, Delalande (MNHN). 1 ♂ Bf 1 - W (SAMC). 1 ♂, 8 unsexed without data (MNHN). Trox instabilis, Haaf 1953: Upon examining the type material of Trox instabilis, we recognised that, according to the current classification, the following nine paratypes are Phoberus sulcatus: 3 ♂♂, 1 ♀ (NHRS: NHRS-JLKB 000073767 - 000073770, aedeagi extracted, examined through detailed photographs): Rectangle, yellowing white, with printed text: “ Cap. B. | Spei. [Cape of Good Hope] || Rectangle, white with handwriting: “ T. sulcatus ” || Rectangle white, printed text interspersed with handwriting: “ Trox ” | instabilis Haaf | det. Dr. E. Haaf 1953 ” || Rectangle, red cardstock with printed text: “ PARATYPE ” || Rectangle white, with handwriting (collector name label): “ Åkerbg. [= Carl-Gustav Åkerberg (1827 – 1888) Swedish Consul in South Africa, donated his collection to Stockholm Museum in 1867] ” || each with different curatorial label, white: “ NHRS-JLKB | 000073767 ”; NHRS-JLKB | 000073768; NHRS-JLKB | 000073769; NHRS-JLKB | 000073770 || Curatorial label, light blue: “ 8656 | E 92 ” || Rectangle, white cardstock with black margin: “ Phoberus sulcatus | (Thunberg, 1787) | det. WP Strümpher 2023 ”. 1 ♀ (TMSA): Rectangle, white with black bars, handwritten text: “ Rietf. | 30.9. [19] 04. ll ” || Narrow rectangle, red: “ PARATYPE ” || Rectangle, white, printed text interspersed with handwriting: “ Trox ” | instabilis Haaf | det. Dr. E. Haaf 1953 ” || Rectangle, white cardstock with orange margin, printed text with handwriting: “ Paratypus 1953 | Trox | instabilis sp. n. | E. Haaf ” || Rectangle, white cardstock with black margin: “ Phoberus sulcatus | (Thunberg, 1787) | det. WP Strümpher 2023 ”. 1 ♂ (MNHN): Rectangle, yellowing white with handwritten text: “ Le Cap ” || Narrow rectangle, red: “ PARATYPE ” || Rectangle, faded blue, printed text: “ MUSEUM PARIS | 1930 | COLL SICARD ” || Rectangle, white cardstock with handwriting: “ Trox | luridus | v. capensis ” || Rectangle, white, printed text interspersed with handwriting: “ Trox ” | instabilis Haaf | det. Dr. E. Haaf 1953 ” || Rectangle, white cardstock with black margin: “ Phoberus sulcatus | (Thunberg, 1787) | det. WP Strümpher 2023 ”. 1 ♀ (SDEI, examined through detailed photographs): Rectangle, yellowing white, handwriting: “ Caffraria ” || Rectangle, yellowing white, with printed text: “ Coll. Kraatz ” || Rectangle, white, handwriting: “ Felsche det ” || Rectangle, white with printed text: “ Dtsch. Entomol. | Institut Berlin ” || Rectangle, yellowing white, printed red text: “ PARATYPUS ” || Rectangle white, printed text interspersed with handwriting: “ Trox ” | instabilis Haaf | det. Dr. E. Haaf 1953 ” || Rectangle white, curatorial label with printed text: “ SDEI Coleoptera | # 303337 ” || Rectangle, white cardstock with black margin: “ Phoberus sulcatus | (Thunberg, 1787) | det. WP Strümpher 2023 ”. 1 ♀ (MNHN): Rectangle, faded white cardstock, handwritten text: “ Cap ” || Narrow rectangle, red: “ PARATYPE ” || Rectangle, faded grey cardstock, printed text: “ MUSEUM PARIS | Coll. L. BEDEL 1922 ” || Rectangle, white, printed text interspersed with handwriting: “ Trox | instabilis Haaf | det. Dr. E. HAAF 1953 ” || Rectangle, white cardstock with black margin: “ Phoberus sulcatus | (Thunberg, 1787) | det. WP Strümpher 2023 ”. 1 ♀ (FMNH, examined through detailed photographs): Rectangle, faded white cardstock, with printed text: “ Cape | Colony ” || white square: “ ♀ ” || Rectangle, white, printed red text interspersed with handwriting: “ PARATYPUS | Trox | instabilis sp. n. | det. Dr. E. HAAF 1953 ” || Rectangle, faded white cardstock, with printed text with handwriting: “ Chicago N. H. Mus [eum]. | ex Museum G. Frey | by exchange 1963 ” || Rectangle white, printed text interspersed with handwriting: “ TROX | sulcatus Thunb. | det. C. H. SCHOLTZ 1979 ” || Rectangle, white cardstock with black margin: “ Phoberus sulcatus | (Thunberg, 1787) | det. WP Strümpher 2023 ”.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF93B5CD7F2FD0AFE55F8FA.taxon	description	Redescription. Size: Length: 9.3 – 13.1 mm. Width: 5.5 – 7.2 mm. Body shape. Convex, elongate oval, elytral margins widest behind the middle, elytral profile convex, broadly rounded, declivous posteriorly, attaining maximum height in the middle. Colour (Figs 1 – 3): Matt black. Body setae mostly yellow, sometimes brownish yellow. Head (Figs 1 – 2): Surface coarsely sculpted with discrete round punctures, punctures irregularly spaced and close, interspaces vary between equal to or less than puncture diameters. Clypeus obtusely triangular, disc slightly elevated, feebly convex, apically deflexed and rounded; anterior clypeal margin entire, bare; clypeogenal and frontoclypeal sutures distinctly raised; frontoclypeal suture complete, with a low clypeogenal tubercle present on either side, tubercles punctate. Frons raised, with paired, arched, setose frontoclypeal carinae delimiting two semicircular depressions, separated by a setose median carina, carinae on frons marked by row of erect, stout seta. Genal angles obtuse; genae wide, shape triangular, surface depressed, coarsely sculpted; genal margin with distinct row of long erect setae. Eyes large and bulbous, scarcely visible in dorsal view. Antennal scape rounded, longer than wide, surface punctate, with long, straight, light brownish yellow setae; pedicel attached apically; antennal club brown, velutinous; mouthparts with dense, yellow to rust-brown setae around. Labrum shape asymmetric; surface of labrum coarsely punctate, each puncture bearing a long stout seta. Pronotum (Figs 1 – 2): Pronotal width only slightly narrower than elytra, widest at base, pronotal length one third of elytral length. Lateral margins broad, explanate and curved, attenuate anteriad, somewhat irregular to coarsely dentate, with fringes of broad spatulate setae; anterior angles acute, posterior angles almost right angled. Base of pronotum marginate (bordered) and projecting backward with weakly produced posteromedian lobe; base obliquely truncate laterally behind posterior angles, entire basal margin with dense fringe of spatulate setae; basal edge with distinct posteromedian notch (opposite scutellum), edge sinuated on either side of notch. Pronotal surface coarsely sculpted with discrete round punctures, irregularly spaced, punctures close, interspaces vary between equal to or less than puncture diameters, surface between punctures dull with superficial fine microreticulation present between punctures. Median discal area of pronotum distinctly raised, broad anteriorly, narrowing towards base forming crudely shaped triangular disc stretching almost over length of pronotum, somewhat flattened anteriorly and strongly declivous on sides and behind; median pronotal ridges distinct, broad anteriorly, with slight depression in the middle, parallel to median groove, ridges interrupted at approximately two-thirds of the length from the anterior margin by a depression; median groove entire stretching almost over length of pronotum, on anterior third widened into slight rounded depression, then partially constricted at approximately two-thirds of the length from the front, thereafter widening into slight round depression (on declivity); median basal tubercles distinct, slightly laterally offset (in line with second elytral costa), not fused to discal ridges; two lateral tubercle (one medially and one basally) present on either side, these tubercle partially fused forming a crude ridge; distinct lateral carina present below lateral tubercles; all tubercles and ridges setose. Scutellum (Fig 1): Longer than wide, oval; surface smooth, apicolaterally rounded and mediobasally depressed; depression weakly sculpted to punctate; margin smooth, tomentose. Elytra (Figs 1 – 2): Elongate, length 2.7 – 2.8 pronotum length, lateral margins subparallel, margin widest at ca. 0.6 length; sides narrow and not explanate, outer lateral margin distinctly dentate or irregular over basal third with setae usually arranged in tufts (if not worn down); setae more regularly distributed over the rest. Humeral calli prominent bearing small tubercles with short, stout setae; humeral angles obtuse, blunt. Sutural margin raised, surface smooth, with low oval to elongate tubercles, widely separated and irregularly distributed along margin, roughly of similar size for 0.75 elytral length, thereafter decreasing in size until barely discernible; these tubercles smooth, dull shiny, each with a posterior setal tuft. Elytral costae distinct, even numbered costae more prominent than odd-numbered. Even-numbered costae with large, raised oval to elongate tubercles, these tubercles smooth, dull shiny, each with a posterior setal tuft; weak apical callosity present on fourth costal interval at top of elytral declivity; costae 2 and 4 with 9 – 11 tubercles for 0.75 elytral length (up to apical callosity), these tubercles close and fairly evenly spaced along elytral length, distance between tubercles roughly equal to or less than length of tubercles; tubercles on costae 6 and 8 more widely spaced, distance between tubercles equal to or greater than length of tubercles. Odd-numbered costae (1, 3, 5, 7, 9) marked by small round to oval, dull shiny tubercles, each with one or two short setae; these tubercles regularly spaced, roughly of equal size for 0.75 elytral length, thereafter decreasing in size until barely discernible. Dorsally costae 1 – 4 separated from intercostae as slightly elevated bands; laterally costae 5 – 9 barely discernible as bands. Intercostae surfaces undulated, marked with distinct round to ovoid punctures (depressions), punctures separated by transverse ridges, punctures evenly spaced along elytral length, distance between ovoid puncture roughly equal to the diameter of puncture; punctures on intercostae roughly equal in size for 0.75 elytral length (up to apical callosity), thereafter slightly decreasing in size; punctures on intercosta 10 (on elytral margin) larger, oval in shape; dorsally margins of intercostae 1 – 4 marked by weak narrow longitudinal ridges (carinae); intercostal carinae laterally vestigial, barely discernible or lacking. Metathoracic wings: Complete. Ventral aspects: Legs (Figs 1 – 3): Protibia dorsally keeled, ending with small upward projecting apical process and row of short stout transverse setae; protibial apex divided into two teeth, distinct in males, females with teeth partly fused; protibial spur stout, pointed forward, flattened laterally, apex slightly curved; as long as, or longer than, third tarsomere; tarsal segments smooth, tarsomeres 1 – 4 subequal in length, fifth tarsomere longest, with two simple and equal claws; tarsomeres 2 – 4 ventrally each with two long seta; outer tibial margin with two distinct median teeth and 4 – 5 smaller basal teeth; in lateral aspect the dorsal longitudinal ridge (keel) on protibia dentate, with row of long, straight setae; dorsally outer tibial surface rugose and punctate, each puncture with short straight seta; both the inner and ventral tibial surfaces smooth and shiny, longitudinally with two crude, shallow, almost continuous parallel grooves (formed from rows of coalesced punctures), grooves on inner tibial surface with numerous long thin setae, setae on ventral surface shorter and less numerous. Procoxa narrow, flattened in front, surface rugose and punctate, basal section indumentose. Profemur large, widest medially, marginate on front face with row of straight forward pointing setae; ventral face with regularly spaced round punctures, each with single short seta; surface dull with superficial reticulation; dorsal surfaces smooth. Meso- and metafemora slender; fore margins rounded, hind margins semi-flattened, marginate with row of stout setae; ventral face rugose, with regularly spaced round punctures, each with single stout seta; dorsal surfaces smooth. Mesotibia and metatibia slender, outer margins dentate and with short, stout yellow setae; inner surface smooth, glabrous, upper and lower margins with long straight setae; outer margin on apex of both meso- and metatibia with row of closely spaced stout backward pointing setae; inner margin of meso- and metatibia with two apical spurs, upper spur slightly longer than lower spur, spurs as long as, or slightly longer than second tarsomere; tarsal segments of both meso- and metatibia smooth, segments 1 – 4 subequal in length, ventrally each with (four to five) long straight backward pointing setae, fifth tarsomere as long as tarsomeres 2 – 4 combined, with two equal and simple claws. Proepisterna (Fig. 3): Surface glabrous, with fine scattered setae, basal margin distinctly raised, marginate; margin punctate with long thin setae; disc with short medial longitudinal ridge. Epipleura: Broad basally narrowing apically; weakly concave along entire length; surface mostly smooth, mid longitudinal row of short setae present along entire length; inner margin distinctly marginate, margin tomentose with scattered short setae. Mesosternum: Sunken below metaventrite, with distinct basolateral depression on either side (visible on clean specimens); surface rugose and coarsely punctate on basal halve, these punctures close, irregular to round or oval, each puncture with single seta; surface on distal halve smooth, sparsely punctate; mesosternum apophysis prominent between mesocoxae, extending almost the length of mesocoxa, narrow (thinly triangular), pointed, with medial longitudinal ridge, surface tomentose. Mesepisternum: surface weakly undulating, rugulose, sparsely setose, distal margin raised, thick. Metepisternum: surface rugulose, with rows of scattered setae along inner margin, inner margin marginate and tomentose. Metasternum: Convex, about 1.7 times as wide as long; surface rugose and coarsely punctate, with evenly scattered short setae; lateral and anterior margin marginate, raised, tomentose; distal margin smooth, plate sunken along mesocoxal margin forming narrow basin like depression; metasternum apophysis prominent between mesocoxae, raised, keeled-like; metaventral disc rhomboidal, visibly depressed medially (discrimen), outer margin with small medial pointed triangular projection; plate strongly declivous behind, transverse edge / margin strongly sinuate on either side of median depression, declivity rugulose and without setae. Abdominal ventrites: Five visible sternites (IV – VIII); sternites IV – VII surface coarsely sculpted with arch-shaped grooves, grooves forming a semicircular pattern, surface between grooves dull shiny, remainder of surface smooth, glabrous, with row of fine punctures running parallel to the distal margin, punctures irregularly spaced, each with short seta, lateral margins setose; distal margin of sternite V straight, subparallel with distal margin of sternite IV, distal margins of sternite VI and VII slightly concave; sternite VIII with distal margin convex, setose, surface finely sculpted, feebly punctate. Male genitalia (Figs 4, 18, 21): Bilaterally symmetrical, slender, strongly sclerotised. Phallobase, elongate, subovate in dorsal view, arched in lateral view; slightly shorter than parameres. Parameres long, slender, extending beyond phallus, in dorsal aspect broadly curved down and inward (Figs 18 a, 21 o); parameres in lateral aspect slender basally, very broadly expanded medially (Fig. 21 n), apically slender, narrowed, apices strongly decurved (never straight or projecting forward) and pointed (Fig. 21). Phallus divided into two symmetrical concave lobes (curved inwards) (Figs 4, 18 b), lobes broad basally, curved sections elevated, inner margins curved (never straight) (Fig, 18 c), apices rounded, flattened, pointed inwards (Fig. 18); two bladelike projections and hooks visible between parameres and phallus.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF93B5CD7F2FD0AFE55F8FA.taxon	diagnosis	Differential diagnosis. The males of Phoberus sulcatus can be distinguished from the males of P. instabilis (Haaf, 1953) and P. erwinhaafi Strümpher, sp. nov. by the following combination of characteristics: the parameres in lateral profile are broad and distinctly enlarged in the middle (Fig. 21 n); the apices are narrow and always curved downward (Fig. 21 m); and the dorsal margin at the apex is always strongly curved or rounded (Fig. 21 o). The three species treated here closely resemble Phoberus cyrtus (Haaf, 1953), Phoberus fumarius (Haaf, 1953), and Phoberus caffer (Harold, 1872), and may be readily confused with them. For detailed morphological descriptions and diagnostic characters that distinguish these taxa, consult Strümpher & Scholtz (2019) and Strümpher & Stals (2021).	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF93B5CD7F2FD0AFE55F8FA.taxon	distribution	Geographic distribution. Phoberus sulcatus is presently known from the Eastern Cape, KwaZulu-Natal, the southeastern part of the Northern Cape and the eastern part of the Western Cape provinces of South Africa (Fig. 24). The distribution of Phoberus sulcatus seems to fall outside of the winter rainfall region of South Africa (roughly east of 21 ° E). Previously, Scholtz (1980), in his monographic revision, reported Phoberus sulcatus (s. l.) occurring as far west as Cape Town, as well as from a few localities from the northwestern part of erstwhile Cape Province [= present-day Northern Cape province] (Scholtz, 1980: 85, Fig. 170). However, during this study, we re-examined all the specimens from Scholtz (1980) that was accessible to us, as well as all other “ sulcatus - like ” specimens collected from west of 21 ° E that have accumulated in South African museums post Scholtz (1980). We found that these specimens were either misidentified or belong to: Phoberus caffer (Harold, 1872) (see Strümpher & Scholtz 2019), Phoberus fumarius (Haaf, 1953) (see Strümpher & Stals 2021), Phoberus luridus (Fabricius, 1782) (see Scholtz 1980) or Phoberus planicollis (Haaf, 1953) (see Scholtz 1980). Even extensive collecting in winter rainfall areas of the Northern and Western Cape provinces over the past two decades yielded no further records of Phoberus sulcatus (WPS personal observation).	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF93B5CD7F2FD0AFE55F8FA.taxon	biology_ecology	Natural history. Collection data show that adults were collected in pitfall traps baited with human dung, carrion (meat), carnivore dung, or a mixture of fresh pig dung and chicken livers. Individuals have also been collected on dried remains of various small and large mammals, such as jackals, bushbucks, cats, cattle, dogs, horses, sheep, vervet monkeys, and rhinos. Based on the collection records, Phoberus sulcatus appears to be active all year round. Immature stages are unknown.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF33B46D7F2FF4AFEA2F8B2.taxon	description	(Figs 6 – 8, 9 – 12, 19, 22, 24)	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF33B46D7F2FF4AFEA2F8B2.taxon	materials_examined	Type locality. Zimbabwe, Harare Metropolitan Province, Harare; as “ Salisbury, Southern Rhodesia ”. Harare is the capital of Zimbabwe and lies at an average elevation of 1,461 m above mean sea level. Type material examined (11 type specimens). Holotype ♂ (NHMB, aedeagus extracted) (Figs 6 – 8): Rectangle, white: “ Salisbury [= Harare; 17 ° 49 ′ S 31 ° 03 ′ E] | S. Rhodesia [Zimbabwe] | Zumpt XI [19] 51 || Rectangle, white: “ Museum Frey | Tutzing ” || Rectangle, white with red bars at top and bottom, printed red text interspersed with black handwriting: “ HOLOTYPUS | Trox | instabilis sp. n. | det. Dr. E. HAAF 1953 ” || Rectangle, white cardstock with black margin: “ Phoberus instabilis | (Haaf, 1953) | stat. rev., comb. nov. | det. Strümpher et al. 2023 ”. Paratype 1 ♂ (NHMB, aedeagus extracted): Rectangle, yellowing white: “ Transvaal | 1883 ” || Narrow rectangle, yellowing white with black bars at top: “ EX MUSEO | N. VAN DE POLL ” || Rectangle, white, printed text interspersed with handwriting: “ PARATYPUS | Trox | instabilis sp. n. | det. Dr. E. HAAF 1953 ” || Rectangle, white cardstock with black margin: “ Phoberus instabilis | (Haaf, 1953) | stat. rev., comb. nov. | det. Strümpher et al. 2023 ”. Paratypes 1 ♂, 1 ♀ (NHMB): Rectangle, yellowing white with printed text: “ Johannesburg [ca. 26 ° 12 ′ S 28 ° 02 ′ E] | Transv. S. Afr. | Zumpt 14 22. XII. [19] 52 ” || Rectangle, white, printed text interspersed with handwriting: “ PARATYPUS | Trox | instabilis sp. n. | det. Dr. E. HAAF 1953 ” || Rectangle, white cardstock with black margin: “ Phoberus instabilis | (Haaf, 1953) | stat. rev., comb. nov. | det. Strümpher et al. 2023 ”. Paratype female with an additional label, white: “ ♀ ”. Paratype 1 ♂ (TMSA, aedeagus extracted): Rectangle, white with green bars on margin, printed text: “ Lydenburg [now Mashishing: South Africa, Mpumalanga, ca. 25 ° 04 ′ S 30 ° 26 ′ E] | Distr. 1896 | P. A. Krantz ” || Narrow rectangle, red: “ PARATYPE ” || Rectangle, white, printed text interspersed with handwriting: “ Trox ” | instabilis Haaf | det. Dr. E. Haaf 1953 ” || Rectangle, white cardstock with orange margin, printed text with handwriting: “ Paratypus 1953 | Trox | instabilis | E. Haaf sp. n. ” || Rectangle, white cardstock with black margin: “ Phoberus instabilis | (Haaf, 1953) | stat. rev., comb. nov. | det. Strümpher et al. 2023 ”. Paratypes 3 ♂♂ (NHMB, aedeagi extracted): Rectangle, faded blue cardstock, printed text: “ MUSEUM PARIS | BASOUTOLAND [Lesotho] | MASERU [Maseru District, 29 ° 18 ′ S 27 ° 29 ′ E] | R. ELLENBERGER 1923 ” || Rectangle, white, printed text interspersed with handwriting: “ PARATYPUS | Trox | instabilis sp. n. | det. Dr. E. HAAF 1953 ” || Rectangle, white cardstock with black margin: “ Phoberus instabilis | (Haaf, 1953) | stat. rev., comb. nov. | det. Strümpher et al. 2023 ”. Paratype 1 ♂ (RBINS, aedeagus extracted, examined through detailed photographs available at https: // virtualcollections. naturalsciences. be / virtual-collections / entomology / coleoptera / trogidae / trox-instabilis-haaf- 1953): Rectangle, faded blue cardstock, printed black text, with two white labels glued to faded blue cardstock, first white label with handwriting, second white label with printed text: Coll. R. I. Sca. N. B. | South Africa || 1 st white label: “ Orange | F. State ” || 2 nd white label: “ EX MUSEO | N VAN DE POLL ” || Rectangle, faded red, printed text framed: “ Para- | type ” || Rectangle white, printed text interspersed with handwriting: “ Trox ” | instabilis Haaf | det. Dr. E. Haaf 1953 ” || Rectangle white, printed text interspersed with handwriting: “ TROX | sulcatus Thunb. | det. C. H. SCHOLTZ 1979 ” || Rectangle, white cardstock with black margin: “ Phoberus instabilis | (Haaf, 1953) | stat. rev., comb. nov. | det. Strümpher et al. 2023 ”. Paratype 1 ♀ (MNHN): Rectangle, faded blue cardstock, printed text: “ MUSEUM PARIS | BASOUTOLAND [Lesotho] | MASERU | R. ELLENBERGER 1923 ” || Narrow rectangle, red: “ PARATYPE ” || white square: “ ♀ ” || Rectangle, white, printed text interspersed with handwriting: “ Trox | instabilis Haaf | det. Dr. E. HAAF 1953 ” || Rectangle, white, printed text interspersed with handwriting: “ TROX | sulcatus Thunb. | det. C. H. SCHOLTZ 1980 ” || Rectangle, white cardstock with black margin: “ Phoberus instabilis | (Haaf, 1953) | stat. rev., comb. nov. | det. Strümpher et al. 2023 ”. Paratype 1 ♀ (MNHN): Rectangle, faded blue cardstock, printed text: “ MUSEUM PARIS | ANCIENNE COLLECTION ” || white square: “ ♀ ” || Narrow rectangle, red: “ PARATYPE ” || Rectangle, white, printed text interspersed with handwriting: “ Trox | instabilis Haaf | det. Dr. E. HAAF 1953 ” || Rectangle, white cardstock with black margin: “ Phoberus instabilis | (Haaf, 1953) | stat. rev., comb. nov. | det. Strümpher et al. 2023 ”. Paratype 1 unsexed (HNHM): Rectangle, yellowing white, printed text with handwriting: “ Africa m. [meridionalis; probably present-day North West province, South Africa] ” | coll. Staudinger ” || Rectangle, yellowing white, printed text with handwriting: Trox | sulcatus | det. J. Frivaldszky || Rectangle, white, printed text with handwriting: Trox | sulcatus | Thoms. | det. Dr. Endrődi || Red cardstock, printed text: “ PARATYPE ” || Rectangle, yellowing white with black border, printed text with handwriting: “ 631 | 12 ” || Rectangle, red framed card, printed text with handwriting: “ Paratypus ” | Trox | instabilis | Haaf ” || Rectangle, white, printed text interspersed with handwriting: “ Trox | instabilis Haaf | det. Dr. E. Haaf 1953 ” || Rectangle, white cardstock with black margin: “ Phoberus instabilis | (Haaf, 1953) | stat. rev., comb. nov. | det. Strümpher et al. 2023 ”.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF33B46D7F2FF4AFEA2F8B2.taxon	discussion	Remarks. Haaf (1953) described Trox instabilis from 23 individuals. We studied the entire type series, which includes the male holotype (NHMB) and 22 paratypes (7 NHMB; 1 FMNH; 2 TMSA; 4 MNHN; 1 RBINS; 4 NHRS; 1 SDEI; 1 NHMW, 1 HMNH). According to the current treatment, the original type series comprises three distinct species. Eleven paratypes belong to Phoberus instabilis revalidated and new combination, nine belong to Phoberus sulcatus (this study), and two belong to the new species described herein.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF33B46D7F2FF4AFEA2F8B2.taxon	materials_examined	Additional material with precise localities examined (502 specimens). ZIMBABWE: Mashonaland Central. 1 ♂ Mazowe [17 ° 31 ′ S 30 ° 58 ′ E], 1895, Darling (SAMC). Matabeleland North. 1 ♂ Victoria Falls, [17 ° 55 ′ S 25 ° 51 ′ E], 10. iv. 1927, RHR Stevenson (SAMC). Bulawayo. 1 ♂ Khami, [20 ° 09 ′ S 28 ° 25 ′ E], 2. xi. 1954 (SAMC). 1 ♀ idem but 25. xi. 1955 (SAMC). 1 ♂ Bulawayo [20 ° 10 ′ S 28 ° 35 ′ E], 18. xi. 1929, RHR Stevenson (TMSA). BOTSWANA: Central District. 1 ♂ Serowe 5 km SE, Farmers Brigade, near Swaneng Hill, [22 ° 24 ′ 44 ″ S 26 ° 45 ′ 32 ″ E], 20. x. 1988, light trap, P Forchhammer (SAMC). 1 ♀ idem but ix. 1989, Malaise trap (SAMC). 1 ♀ Serowe 4 km S, Swaneng Hill School, [22 ° 24 ′ 47 ″ S 26 ° 44 ′ 54 ″ E], mercury vapour lamp, P Forchhammer (CMNC). SOUTH AFRICA: Limpopo. 1 ♀ Soutpansberg, Ingwe Motel [now Ingwe Ranch], 22 ° 58 ′ S 29 ° 56 ′ E, 1,200 m, 23 – 24. xi. 1992, H Hölzel, P Ohm, MW Mansell (SANC). 1 ♂ Soutpansberg, near Vivo, Goro Research Camp, 22 ° 59 ′ S 29 ° 26 ′ E, 19. iii. 2018, light trap, WP Strümpher (TMSA). 1 ♀ Soutpansberg, Lajuma Research Centre, [23 ° 02 ′ S 29 ° 26 ′ 30 ″ E], 16. i. 2016, C Munyai, P Tshililo (TMSA). 2 ♂, 3 ♀ Soutpansberg, Schoemansdal [environmental education centre], [ca. 23 ° 01 ′ S 29 ° 43 ′ E], 2. v. 1977, CH Scholtz (SAMC). 1 spec. unsexed idem (CMNC). 1 spec. unsexed idem (RMCA). 1 ♂ Louis Trichardt [23 ° 03 ′ S 29 ° 54 ′ E], 6. ii. 1952 (SAMC). 3 ♂, 3 ♀ Vivo 21 km N, Blouberg Nature Reserve, 23 ° 01 ′ S 29 ° 04 ′ E, 991 m, 20. xii. 2003, A Frolov, C Deschodt (TMSA). 2 unsexed Blouberg at Blouberg settlement, 23 ° 05 ′ S 28 ° 59 ′ E, 1,680 m, 4 – 7. xii. 1990, SL Chown, HE Steenkamp, MA McGeogh (SANC). 1 ♂ Polokwane, [23 ° 54 ′ S 29 ° 27 ′ E], JP Cregoe (SAMC). 1 ♂ Mookgophong, Bundy Lodge, 24 ° 27 ′ S 28 ° 42 ′ E, 29. x. 2004, WP Strümpher (SAMC). 1 ♂ Waterberg, Meletse [farm], 24 ° 36 ′ S 27 ° 39 ′ E, 13. ii. 2016, R Müller (TMSA). 1 ♂ Thabazimbi, Ben Alberts Nature Reserve, 24 ° 37 ′ S 27 ° 23 ′ E, 24 – 28. xi. 1980, SJ van Tonder (SANC). 1 ♂ Mookgophong, Nylsvley Nature Reserve, 24 ° 39 ′ S 28 ° 42 ′ E, 28. xi. 1979, E Grei (SANC). 1 ♀ idem but ix. 1980 (SANC). 1 ♂, 1 ♀ Radium, Langkuil Private Game Farm, 20 km S Warmbad; ca. 25 ° 02 ′ S 28 ° 11 ′ E], T Beyers (TMSA). Mpumalanga. 1 ♂ Kruger National Park, Skukuza Rest Camp [24 ° 59 ′ S 31 ° 36 ′ E], xii. 1977, CH Scholtz (SAMC). 1 ♀ Siyabuswa, 25 ° 07 ′ S 29 ° 03 ′ E, 21. ii. 1998, Poto (TMSA). 1 ♀ Roossenekal, Booysensdal Mine North, 25 ° 07 ′ S 30 ° 08 ′ E, 25. x. 2000, E-Y: 3435, at light, Museum staff (TMSA). 1 ♂ Stofberg, Laersdrif, [25 ° 22 ′ S 29 ° 51 ′ E], 8 – 13. xi. 1974, J Dedman (SAMC). 1 ♂ Mbombela, De Hoop 203 [farm], [ca. 25 ° 32 ′ S 31 ° 09 ′ E], 20 – 21. xi. 1972, A Strydom (TMSA). 1 ♀ Barberton, Saddleback Hill [ca. 25 ° 47 ′ S 31 ° 05 ′ E], Barberton, xii. 1998, van Viegen (TMSA). 7 ♂ Lake Chrissie farm, 5 km SE of Chrissiesmeer, 26 ° 18 ′ S 30 ° 15 ′ E, 1,700 m, 26. xi. 2002, in remains of dead sheep, R Stals (SANC). Gauteng. 1 ♂ Bronkhorstpruit, Ezemvelo Nature Reserve, 25 ° 42 ′ S 29 ° 01 ′ E, 1,374 m, 26. i. 2004, E-Y: 3605, light trap, TMSA staff (TMSA). 1 ♂ idem but 27. i. 2004, E-Y: 3606, pitfall trap, R Müller (TMSA). 1 ♀ Bronkhorstpruit, Rhenosterpoort Private Nature Reserve, 25 ° 45 ′ S 28 ° 58 ′ E, 19. ii. 1975, PS Pots & MJ Scoble (TMSA). 2 ♂, 1 ♀ idem but 16. xi. 1975, E-Y: 937, bushveld, sifted from litter, S Endrödy-Younga (TMSA). 1 ♀ Boekenhoutskloof [farm], 25 ° 31 ′ S 28 ° 26 ′ E, 25. iii. 1994, J Harrison (TMSA). 2 ♀ Pretoria 25 km E, Bashewa Plot 12, 25 ° 56 ′ S 28 ° 24 ′ E, 1,550 m, 28. xii. 1995, R Stals (SANC). 1 unsexed Pretoria, Roodeplaat [ARC experimental farm], 25 ° 36 ′ S 28 ° 21 ′ E, 22. xi. 1979, M Bolton (SANC). 1 ♂ idem but x. 1979, C Kok (SANC). 1 ♀ idem but xi. 1979, SJ van Tonder (SANC). 1 ♀ 1 ♂ Pretoria, Shere [25 ° 48 ′ S 28 ° 21 ′ E], 27. x. 1963, R Hofman (TMSA). 1 ♀ Pretoria, [ca. 25 ° 44 ′ S 28 ° 13 ′ E], xi. 1924, HJ Heske (TMSA). 11 ♂ idem but 5. iii. 1931, C. S. (SAMC). ♂ idem but 5. iv. 1932, DHB (SAMC). 1 ♀ Pretoria, 1.1.1975, L Schulze (TMSA). 1 ♀ Pretoria North, 25 ° 41 ′ S 28 ° 11 ′ E, ix. 1937, G van Son (TMSA). 4 ♂, 2 ♀ Pretoria, Onderstepoort Nature Reserve, Kaalplaas, [25 ° 38 ′ S 28 ° 10 ′ E], 11 – 16. v. 1977, J Boomker (SAMC). 2 spec. unsexed idem but 16. v. 1977 (RMCA). 2 ♂, 1 ♀ Broederstroom, Vlakplaas portion 53 [farm], Protea Ranch, [25 ° 49 ′ 30 ″ S 28 ° 01 ′ E], 2 – 15. xi. 2008, M Paulsen (TMSA). 1 ♂ Johannesburg, [ca. 26 ° 12 ′ S 28 ° 02 ′ E], x. 1901 (TMSA). 1 ♂ idem but xii. 1934, G Kobrow (SAMC). 1 ♂ idem but 30. xii. 1945, A Capener (TMSA). 1 ♀ idem but iii. 1963, F Zumpt (NHMB). 1 ♀ Johannesburg, Klipriviersberg Municipal Nature Reserve [26 ° 18 ′ S 28 ° 01 ′ E], 3. xi. 2013 (TMSA). 1 ♂, 1 ♀ West Rand District Municipality, Nascott Game Park [now John Nash Nature Reserve,], 25 ° 54 ′ S 27 ° 46 ′ E, pitfall traps: 1. xi. 1985, E-Y: 2567, G du Plessis (TMSA). 1 ♀ idem but 28. xi. 1985, E-Y: 2568 (TMSA). 1 ♀ idem but 3. iii. 1986, E-Y: 2571 (TMSA). 2 ♀ idem but 17. iv. 1986, E-Y: 2572 (TMSA). 1 ♂ idem but 14. viii. 1986, E-Y: 2574 (TMSA). 1 ♂ idem but 1. xi – 29. xi. 1985 (TMSA). 1 ♂ idem but 23. ix – 23. xi. 1986 (TMSA). 1 ♂ Krugersdorp Game Reserve [26 ° 06 ′ S 27 ° 43 ′ E], 30. ix. 2008, light trap, E Pretorius (TMSA). 1 ♂, 2 ♀ Vereeniging, [26 ° 40 ′ S 27 ° 56 ′ E], 15. xii. 1915 (SANC). North West. Hartbeespoort, Daspoortrant, Saartjiesnek [25 ° 46 ′ S 27 ° 56 ′ E], ii. 1978, E Holm (SAMC). 1 ♂ Hartbeespoort, [ca. 25 ° 44 ′ S 27 ° 53 ′ E], i. 1961, E Holm (SAMC). 1 ♀ Rustenburg, Buffelspoort Dam, 25 ° 47 ′ S 27 ° 29 ′ E, 9. x. 1983, RG Oberprieler (SANC). 1 ♀ Rustenburg Nature Reserve, 25 ° 40 ′ S 27 ° 12 ′ E, 17 – 20. iii. 1980, WA Harrop (SANC). 2 ♀ Venterskroon, Vredefort Dome, 26 ° 50 ′ S 27 ° 19 ′ E, 1,388 m, 19 – 20. ii. 2012, E-Y: 3934, disturbed habitat, R Müller (TMSA). 1 ♂, 1 ♀ idem but E-Y: 3935, at light (TMSA). 1 ♀ Potchefstroom [ca. 26 ° 43 ′ S 27 ° 05 ′ E], 28. xii. 1976, TW Drinkwater (SANC). 1 ♂ idem but 7. i. 1977 (SAMC). 1 ♂ idem but 5. iii. 1988, AE Gildenhuys (SAMC). 1 ♀ Koster, 25 ° 52 ′ S 26 ° 54 ′ E, 16. xi 1993, R Stals, M Vogt, F Joubert (SANC). 1 ♂ Zeerust 23 km NE, Meseg, Enzelsberg, 25 ° 22 ′ S 26 ° 14 ′ E, 20 – 22. ii. 2009, 1,300 m, CD Theron (CEMT). Northern Cape. 1 ♂ Jan Kempdorp, [ca. 27 ° 55 ′ S 24 ° 50 ′ E], i. 1961, H. S. (SAMC). 2 ♂, 1 ♀ Barkly West, Sydney on Vaal, 28 ° 27 ′ S 24 ° 19 ′ E, 7 – 9. iii. 1990, S Louw (BMSA). 1 ♂ Barkly West, Vaalbos National Park [deproclaimed], 28 ° 34 ′ S 24 ° 18 ′ E, 7 – 9. iii. 1990, S Louw (BMSA). 1 ♀ Kimberley, [ca. 28 ° 43 ′ S 24 ° 45 ′ E], 30. i. 1948, RJ Power (SAMC). 1 ♂ idem but 3. iv. 1948 (SAMC). 1 ♀ Kimberley, Dronfield Nature Reserve [ca. 28 ° 37 ′ S 24 ° 48 ′ E]. 1. iv. 2023, H de Klerk (TMSA). 1 ♀ Kimberley, Riverton [28 ° 31 ′ S 24 ° 42 ′ E], 16. xi. 1961, HAD van Schalkwyk (SANC). 1 ♀ Kimberley, Langeberg 138 [farm], 28 ° 55 ′ S 24 ° 36 ′ E, pitfall traps: xi. 1987 – iii. 1988 (BMSA). 1 ♂, 2 ♀ Schmidtsdrif, Rooipoort [farm], 28 ° 38 ′ S 24 ° 16 ′ E, 1,129 m, 3. iii. 2009, E-Y: 3847, Kalahari bushveld, light trap, D MacFayden, R Müller (TMSA). 1 ♀ Schmidtsdrif, [28 ° 42 ′ S 24 ° 03 ′ E], 2. x. 1954, G van Son (TMSA). 1 ♂ Kuruman [27 ° 28 ′ S 23 ° 25 ′ E], 8. iv. 1933, G van Son (TMSA). 1 ♀ Sishen 5 km S, 27 ° 50 ′ S 22 ° 59 ′ E, xi. 1996, light at night, E Holm (SANC). 1 ♀ Groblershoop, 28 ° 53 ′ S 21 ° 59 ′ E, 1 – 7. iv. 1980, CG Moolman (SANC). Free State. 1 ♀ Sasolburg, 26 ° 49 ′ S 27 ° 49 ′ E, 20. xi. 1988, N Duke (SANC). 1 ♂ Parys 4 km E, Abel 52 [farm], 26 ° 54 ′ S 27 ° 35 ′ E, 4 – 7. xii. 1992, M Krüger (TMSA). 1 ♂ Koppies, [27 ° 14 ′ S 27 ° 35 ′ E], 1. i. 1982, RS van Zyl (BMSA). 1 unsexed Heilbron, Uitsien 1449 [farm], SE 2727 Db, 26 – 30. xii. 1982, A van Rensburg (BMSA). 1 ♀ Bothaville, [27 ° 24 ′ S 26 ° 37 ′ E], 30. iv. [18] 99, Dr Brauns (TMSA). 4 ♂, 2 ♀ Bothaville, Deelfontein 482 [farm], 27 ° 07 ′ S 26 ° 35 ′ E, 12 – 22. iv. 1977, GW & MC Ferreira (BMSA). 1 ♂, 1 ♀ idem but 13 – 17. ix. 1978, A Strydom (BMSA). 10 ♂, 16 ♀ idem but viii – x. 1986 (BMSA). 1 ♂ Kroonstad [27 ° 40 ′ S 27 ° 14 ′ E], 17. i. 1965, DJ Brothers (TMSA). 1 spec. unsexed Lindley, Middelpunt 760 [farm], 28 ° 05 ′ S 28 ° 00 ′ E, ix. 1985, A van Wyk (BMSA). 1 ♂ Hoopstad, Sandveld Nature Reserve, [ca. 27 ° 38 ′ S 25 ° 41 ′ E], 8 – 12. ii. 1982, Entomology Dept. (BMSA). 1 ♀ Hoopstad, Swartsrus 1193 [farm], 27 ° 45 ′ S 25 ° 30 ′ E, iv. 1985, Entomology Dept. (BMSA). 6 ♂, 2 ♀ Hoopstad, Sophiasdal 280 [farm], [ca. 27 ° 54 ′ S 25 ° 45 ′ E], 7 – 8. ii. 1978, A Strydom (BMSA). 1 ♀ idem but 4. iii. 1978 (BMSA). 1 ♀ idem but iii – v. 1982 (BMSA). 1 ♂ Bultfontein, Hartenbosch 43 [farm], [ca. 28 ° 17 ′ S 26 ° 01 ′ E], 1 – 2. ii. 1978, A Strydom (BMSA). 3 ♂, 2 ♀ Boshof, [28 ° 32 ′ S 25 ° 14 ′ E], 4. xii. 1976, E Holm (SAMC). 1 ♂ idem (RMCA). 1 spec. unsexed idem but CH Scholtz (CMNC). 1 spec. unsexed Boshof 20 km W, [ca. 28 ° 32 ′ S 25 ° 01 ′ E], 12 – 14. xii. 1984, H Howden (CMNC). 9 ♂, 9 ♀ Voorspoed 1585 [farm], near Boshof, SE 2825 Cd [ca. 28 ° 52 ′ S 25 ° 46 ′ E], 21 – 28. iii. 1977, A Strydom (BMSA). 1 ♀ idem but 15 – 16. ii. 1978 (BMSA). 2 ♂, 2 ♀ Boshof, Wesselsdam 379 [farm], near SE 2825 Ba [ca. 28 ° 09 ′ S 25 ° 36 ′ E], 15 – 16. ii. 1978, A Strydom (BMSA). 1 ♀ idem but 21 – 28. iii. 1977 (BMSA). 1 ♀ Winburg [ca. 28 ° 31 ′ S 27 ° 01 ′ E], v. [19] 07, H Fry (TMSA). 6 ♂, 4 ♀ Vrede, Beginsel 1284 [farm], Vrede, [ca. 27 ° 26 ′ S 28 ° 44 ′ E], 9 – 15. ii. 1979, A Strydom (BMSA). 1 ♀ Harrismith, Greenlands 1245 [farm], 28 ° 16 ′ S 28 ° 59 ′ E, 17 – 17. xii. 1984, A van Rensburg (BMSA). 4 ♂, 1 ♀ Harrismith, Kerkenberg, [28 ° 30 ′ S 29 ° 08 ′ E], 1 – 12. ii. 1977, GW & MC Ferreira, A Strydom (BMSA). 38 ♂, 30 ♀ 126 spec. unsexed Ladybrand, Nova 667 [farm], [29 ° 07 ′ S 27 ° 22 ′ E], 1 – 12. ii. 1977, GW & MC Ferreira, A Strydom (BMSA). 1 ♂, 1 ♀ idem but 19 – 22. i. 1976, GW Ferreira & Ee (BMSA). 1 ♂ idem but 15 – 26. v. 1978, GW Ferreira & A Strydom (BMSA). 1 ♂, 3 ♀ Mowbray 287 [farm], near Ladybrand, SE 2927 Ab, 18 – 29. ix. 1978, Museum staff (BMSA). 4 ♂, 6 ♀ Groenkloof 294 [farm], near Ladybrand, SE 2927 Ab, 1 – 12. ii. 1977, GW & MC Ferreira (BMSA). 1 ♂, 1 ♀ idem but 18 – 29. ix. 1978, Museum staff (BMSA). 1 ♂, 1 ♀ Winnie Mandela, Florisbad 686 [farm, research station] [28 ° 46 ′ S 26 ° 04 ′ E], i. 1984, Entomology Dept. (BMSA). 1 ♀ idem but xii. 1984 (BMSA). 1 ♀ idem but iv. 1985 (BMSA). 2 ♂, 1 ♀ idem but ix. 1985 (BMSA). 2 ♀ idem but x. 1985 (BMSA). 2 spec. unsexed idem but 9 – 12. xii. 1984, H & A Howden (CMNC). 5 ♂, 6 ♀ Bloemfontein, Glen [farm, agricultural college], 28 ° 56 ′ S 26 ° 20 ′ E, 18. x. 1983, Van Rensburg & Earlé (BMSA). 2 ♂, 3 ♀ idem but 22 – 30. iii. 1977, A Strydom (BMSA). 4 ♂, 4 ♀ idem but 5. iv. 1978 (BMSA). 2 spec. unsexed Bloemfontein, [ca. 29 ° 07 ′ S 26 ° 13 ′ E], 15. ii – 14. iii. 1976, Museum staff (BMSA). 1 ♂, 1 ♀ Bloemfontein, Free State National Botanical Garden, [ca. 29 ° 03 ′ S 26 ° 13 ′ E], ii. 2006, Museum staff (BMSA). 2 ♂ Bloemfontein, Monument 2206 [farm], SE 2926 Ac, 7. xii. 1976, A Strydom (BMSA). 1 ♂ idem but 17 – 18. viii. 1983 (BMSA). 4 ♂, 5 ♀ Bloemfontein, Merino 14 [farm], [ca. 28 ° 52 ′ S 26 ° 12 ′ E], 29. xi – 3. xii. 1976, A Strydom (BMSA). 1 ♀ Bloemfontein, The Willows 5837 [farm], 29 ° 14 ′ S 26 ° 05 ′ E, 8. ix. 1983, Van Rensburg & Earlé (BMSA). 1 ♂, 1 ♀ Fauresmith, Zuurfontein 92 [farm], [ca. 29 ° 55 ′ S 25 ° 24 ′ E], 19 – 22. i. 1982, JH van Wyk (BMSA). 1 ♂, 1 ♀ Philippolis, Allemans Drift 37 [farm], [ca. 30 ° 32 ′ S 25 ° 20 ′ E], 23 – 29. x. 1978, A Strydom (BMSA).	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF33B46D7F2FF4AFEA2F8B2.taxon	description	Redescription. Size: Length: 9 – 12.5 mm. Width across humeri: 5.6 – 7.3 mm. Body shape: Elongate oval, elytral margins subparallel, elytral profile somewhat flat to slightly convex, attaining maximum height in middle, declivous posteriorly. Colour (Figs 9 – 10): Matt black. Body setae mostly brownish yellow. Head (Figs 9 – 10): As in Phoberus sulcatus. Pronotum (Figs 9 – 10): As in Phoberus sulcatus, except pronotal tubercles and ridges are higher and more prominent, punctation appears denser, pronotal surface slightly more embossed. Scutellum (Fig. 9): Longer than wide, obround in shape; surface smooth; medio-basally depressed, depression punctate; margin smooth, tomentose. Elytra (Figs 9 – 10): Relative dimensions and margins as in Phoberus sulcatus, except that the profile is somewhat flatter. Humeral calli, and sutural margin as in Phoberus sulcatus. Elytral costae distinct, even numbered costae more prominent than odd-numbered. Even-numbered costae with large raised oval to elongate tubercles, these tubercles smooth, dull shiny, each with a posterior setal tuft; tubercles on costae 2 and 4 irregularly spaced along elytral length, distance between tubercles roughly equal to or more than length of tubercles; tubercles on costae 6 and 8 widely spaced, distance between tubercles equal to or greater than length of tubercles; weak apical callosity present on fourth costal interval at top of elytral declivity. Elytral costae 2 and 4 with 7 – 9 tubercles for 0.75 elytral length (up to apical callosity). Odd-numbered costae as in Phoberus sulcatus. Dorsally costae 1 – 4 separated from intercostae as slightly elevated bands; laterally costae 5 – 9 barely discernible as bands. Intercostae as in Phoberus sulcatus. Metathoracic wings: Complete. Ventral aspects: As in Phoberus sulcatus. Male genitalia (Figs 7, 11, 19, 22): Symmetrical, slender, strongly sclerotised. Phallobase, subovate in dorsal view, arched in lateral view, as long as or slightly shorter than parameres. In dorsal aspect parameres long, slender, extending well beyond phallus, attenuated inward, apex narrowed, pointed, always projecting forward (never inward or down) (Figs 11, 19 d). Parameres in lateral aspect evenly broad basally, sides subparallel, feebly sinuate apically (Fig. 22 r; never convex or curved as in Fig. 21 o), moderately expanded medially (compare Fig. 22 q with Fig. 21 n), apically slender, narrowed, apices long, only moderately decurved (almost straight, never strongly decurved or projecting downward as in Fig 21 m), projecting forward (Figs 22 p). Phallus divided into two symmetrical lobes (Figs 11, 19); lobes broad basally, inner halve of lobe there elevated, concave (Fig. 19), lobes strongly narrowing in apical third, outside margin notched at apical third (Fig. 19 f), apex of phallus narrow, apex broadly rounded and pointing inward, weakly keeled (Fig. 19 e); inner margins of lobes almost parallel (Fig. 19 g); two bladelike projections and hooks visible between parameres and phallus.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF33B46D7F2FF4AFEA2F8B2.taxon	diagnosis	Differential diagnosis. Phoberus instabilis revalidated and new combination is the most similar to Phoberus sulcatus, but males can be distinguished from the latter by the following combination of characteristics: the parameres in lateral profile are broad, weakly expanded medially (Fig. 22 q); their apices are long, narrow and weakly curved downward (Fig. 22 p); and the dorsal margin at the apex is primarily straight and feebly concave (Fig. 22 r). Furthermore, in Phoberus instabilis the lobes on the phallus are broad and not strongly concave as in Phoberus sulcatus (compare Figs 18 and 19). Also, see Table 1 for additional characteristics.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF33B46D7F2FF4AFEA2F8B2.taxon	distribution	Geographic distribution. Phoberus instabilis is known from Botswana, Lesotho, South Africa and Zimbabwe (Fig. 24). In South Africa, the species has been recorded from the Free State, Gauteng, Limpopo, Mpumalanga, North West and Northern Cape provinces.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFF33B46D7F2FF4AFEA2F8B2.taxon	biology_ecology	Natural history. Based on label data, adult beetles have been collected at light in the night and from the remains of a dead sheep. Collection records show Phoberus instabilis to be active from August to May. Immature stages are unknown.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFEE3B4ED7F2F891FB34FEAA.taxon	description	(Figs 13 – 17, 20, 23, 24)	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFEE3B4ED7F2F891FB34FEAA.taxon	materials_examined	Type locality. Namibia, Khomas Region, Windhoek c. 10 km WSW, Regenstein Farm, 22 ° 36 ′ S 16 ° 59 ′ E; verbatim as in holotype label data below. Located on the central plateau of Namibia, the approximate elevation of this farm is 1,700 m above mean sea level. In the pertinent collecting notebook of Sebastian Endrödy-Younga, collector of the holotype, this locality is annotated with “ Farm de Wet ”. We could not trace a farm called “ De Wet ” in Namibia, hence we assume that De Wet, a fairly common Afrikaans surname, was the surname of the owner of, or farmer on, the farm Regenstein when part of the type series was collected there. Type material (28 type specimens). NAMIBIA: Khomas Region. Holotype ♂ (TMSA, specimen cleaned, aedeagus extracted) (Figs 13 – 17): Rectangle white, printed text: “ S. W. Afr. [South West Africa, now Namibia], Windhoek | Regenstein | 22.36 S- 16.59 E [22 ° 36 ′ S 16 ° 59 ′ E] ” || Rectangle white, printed text: “ 21.1 [January]. 1975, E-Y: 562 | groundtrap [pitfall trap], 44 day [s] | leg. Endrödy-Younga ” || Rectangle white, printed text: “ ground traps | with meat bait ” || Rectangle white, printed text with handwriting: “ TROX | sulcatus Thunb. | det. C. H. SCHOLTZ 1979 ” || Rectangle, red cardstock, preprinted black text: “ HOLOTYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ” || Rectangle white, printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000864 ”. The following specimens are designated as paratypes, representing individuals collected across the distribution range of the new species. During the examination of the type series of Trox instabilis Haaf, 1953, we identified two specimens (housed in NHMB and NHMW) that match the diagnostic features of the new species; these are included as paratypes. All other examined material (35 specimens), though not part of the paratype series, is herein referred to as “ additional specimens ”. NAMIBIA: Oshana Region. Paratype 1 ♂ (TMSA): Rectangle, white, printed text: “ S. Afr., Etosha [Etosha National Park] | Okaukuejo [Rest] Camp [19 ° 11 ′ S 15 ° 55 ′ E] | Dixon, 29. II [February]. 1976 ” || Rectangle, white, printed text with handwriting: “ Trox | sulcatus Thunb | det. C. H. Scholtz 1982 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ” || Rectangle white, printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000925 ”. Otjozondjupa Region. Paratype 1 ♂ (NHMW NHMW-ZOO-COL- 0000154): Rectangle, yellowing white, printed text: “ Farm Okonjati | Okowakuatjiwi ” || Rectangle, yellowing white, printed text: “ D. S. W. Afrika [Namibia] | J. Schneider ’ 09 ” || Rectangle, red cardstock, printed text: “ PARATYPE ” || Rectangle, white, printed text: “ Coll. Mus. | Vindob. ” || Rectangle, white, printed text interspersed with handwriting: “ Trox | instabilis Haaf | det. Dr. E. Haaf 1953 ” || Rectangle, white, with handwriting: “ Trox sulcatus | Thunberg | Det. Pittino 1989 ” || Rectangle, white, printed text, curatorial label: “ NHMW-ZOO- | COL- 0000154 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ”. Erongo Region. Paratype 1 ♂ (TMSA): Rectangle, yellowing white, printed text: “ ERONGO Mtns. [ca. 21 ° 42 ′ S 15 ° 24 ′ E] | Namib side [western slopes] | III [March]. 1956 | W. Hoesch ” || Rectangle, yellowing white, printed text with handwriting: “ Trox | instabilis M. [male] | det. Dr. Haaf, 1957 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ” || Rectangle white, printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000926 ”. Khomas Region. Paratypes 5 ♂, 1 ♀ (TMSA): Labels identical to those of holotype but with different determination and database labels: “ PARATYPE: | Phoberus erwinhaafi | det. WP Strümpher 2023 ” || Rectangle white, printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000876 [plus WTD 0000877 to WTD 00000881] ”. Paratypes 2 ♂, 1 ♀ (TMSA): Rectangle white, printed text: “ S. W. Afr. [South West Africa, now Namibia], Windhoek | Regenstein | 22.36 S- 16.59 E [22 ° 36 ′ S 16 ° 59 ′ E] ” || Rectangle white, printed text: “ 3.12.1974, E-Y: 499 | groundtrap [pitfall trap], 48 days | leg. Endrödy-Younga ” || Rectangle white, printed text: “ ground traps | with meat bait ” || Rectangle white, printed text with handwriting: “ TROX | sulcatus Thunb. | det. C. H. SCHOLTZ 1979 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ” || Rectangle white, printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000873 [plus WTD 0000874 to WTD 0000875] ”. Paratypes 2 ♂, 1 ♀ (TMSA): Rectangle white, printed text: “ S. W. Afr. [South West Africa, now Namibia], Windhoek | Regenstein | 22.36 S- 16.59 E [22 ° 36 ′ S 16 ° 59 ′ E] ” || Rectangle white, printed text: “ 7.3 [March]. 1974; E-Y: 724 | groundtrap [pitfall trap], 185 day [s] | Endrödy [- Younga] & Breytenb [ach]. ” || Rectangle white, printed text: “ ground traps | with meat bait ” || Rectangle white, printed text with handwriting: “ TROX | sulcatus Thunb. | det. C. H. SCHOLTZ 1982 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ” || Rectangle white, printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000870 [plus WTD 0000871 to WTD 0000872] ”. Paratype 1 ♂ (CJBH): Rectangle white, printed text: “ NAMIBIA, Khomas Mts. [Mountains], | 35 km W Windhoek, | 22 ° 34 ′ 46 ″ S, 16 ° 45 ′ 41 ″ E, | 1800 m, 4 – 9. I. 2019, G. Betti lg. ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ”. Paratype 1 ♂ (NHMB): Square, yellowing white: “ S. W. AFRICA; | Hoffnung. | 1,850 m. | 5. i. 1934. | K. Jordan. ” || Narrow rectangle, yellowing white: “ Brit. Mus. | 1934 - 288. ” || Rectangle, white, printed text interspersed with handwriting: “ PARATYPUS | Trox | instabilis sp. n. | det. Dr. E. HAAF 1953 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ”. Paratype 1 ♀ (TMSA): Rectangle, yellowing white, printed text: “ Gocheganas 26 [farm, nature reserve] | 22 ° 50 ′ S., 17 ° 11 ′ E | WINDHOEK DISTRICT | 22 Feb. – 20 March 1982 | Preserv. traps | M. - L. Penrith ” || Rectangle, white, printed text: “ H 56959 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ” || Rectangle white, printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000927 ”. Omaheke Region. Paratype 1 ♀ (SDEI, examined through detailed photographs): Rectangle white, handwriting: “ Gobabis ” [22 ° 27 ′ S 18 ° 58 ′ E] || Rectangle white, printed text: “ Coll. v. Bennigsen ” || Rectangle white, handwriting: “ Kolbe Oct. ” || Rectangle white, printed text: “ Dtsch. Entomol. | Institut Berlin ” || Rectangle white, printed text with handwriting: “ Trox | sulcatus Thunb. | det. Dr. E. Haaf 1953 ” || Rectangle white, printed text: “ SDEI Coleoptera | # 303410 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ”. Hardap Region. Paratypes 1 ♂, 2 ♀ (TMSA): Rectangle white, printed text: “ S. W. Afr., Nauwkluft | Nauwkluft Park | 24.16 S- 16.15 E [24 ° 16 ′ S 16 ° 15 ′ E] ” || Rectangle white, printed text: “ 26.10.1974 | groundtraps [pitfall traps] | leg. Endrödy-Younga ” || Rectangle white, printed text: “ groundtrap | with faeces bait ” || Rectangle white, printed text with handwriting: “ TROX | sulcatus Thunb. | det. C. H. Scholtz 1979 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ” || Rectangle white, printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000928 [plus WTD 0000929 to WTD 0000930] ”. Karas Region. Paratypes 2 ♂, 1 ♀ (CJBH): Rectangle white, printed text: “ Namibia, Karas Region, | Savanna Guest Farm, | 35 km N of Grünau, 1450 m, | 27 ° 22 ′ 57 ″ S, 18 ° 31 ′ 11 ″ E, | 19. II. 2012, leg. M. Hornburg ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ”. SOUTH AFRICA: Northern Cape Province: Kgalagadi Transfrontier Park. Paratype 1 ♂ (TMSA): Rectangle white, printed text: “ S. Afr., Kalh. Park | Mata Mata | 25.45 S- 20.00 E [25 ° 45 ′ S 20 ° 00 ′ E] ” || Rectangle white, printed text: “ 18.12.1974; E-Y: 512 | at light | leg. Endrödy-Younga ” || Rectangle white, printed text with handwriting: “ Trox | sulcatus Thunb | det. C. H. Scholtz 1982 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ” || Rectangle white, printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000931 ”. Paratype 1 ♂ (TMSA): Rectangle, yellowing white, printed text with handwriting: “ Twee Rivieren [Rest Camp, 26 ° 28 ′ S 20 ° 37 ′ E] | 12 – 15 April | det. C. H. Scholtz ’ [19] 89 ” || Rectangle, red cardstock, preprinted black text: “ PARATYPE | Phoberus erwinhaafi | det. WP Strümpher 2023 ” || Rectangle white, printed text: “ [QR Code] | WORLD | TROGIDAE | DATABASE | WTD 0000932 ”. Additional material with exact localities (35 specimens). NAMIBIA: Oshana Region. 1 ♂, 4 ♀ Etosha National Park, Okaukuejo Rest Camp, [19 ° 11 ′ S 15 ° 55 ′ E], i. 1978, RG Oberprieler (SAMC). Erongo Region. 1 ♂ Omatjete, near Omaruru [21 ° 03 ′ S 15 ° 30 ′ E], xii. 1966, E Kölle (TMSA). Khomas Region. 1 ♂, 1 ♀ Windhoek, [22 ° 34 ′ S 17 ° 05 ′ E], Nagel (RMCA). 1 ♀ Excelsior 286 [farm], Seeis, Windhoek ENE, [ca. 22 ° 27 ′ S 17 ° 38 ′ E], iv. 1978 (SAMC). 1 ♀ Hoffnung, Windhoek E, [22 ° 33 ′ S 17 ° 12 ′ E], 1,850 m, 5. i. 1934, K Jordaan (NHMB). 2 ♀ Gocheganas 26 [farm, nature reserve], Windhoek S, 22 ° 50 ′ S 17 ° 09 ′ E, 22. ii – 20. iii. 1982, M-L Penrith (SAMC). 1 ♂, 1 ♀ Isabis 19 [farm], Windhoek SW, 22 ° 58 ′ S 16 ° 46 ′ E, 24 – 25. xii. 1974 (SAMC). 1 ♂ Hohenheim farm, Khomashochland, 23 ° 18 ′ S 16 ° 23 ′ E, pitfall traps: 8. iii. 1975 - 4. ix. 1975, E-Y: 729, S Endrödy-Younga (TMSA). 1 ♀ Weissenfels farm, Khomashochland, 23 ° 18 ′ S 16 ° 27 ′ E, pitfall traps: 21. x. 1974 - 6. xi. 1974, E-Y: 403, S Endrödy-Younga (TMSA). 2 ♂, 1 ♀ idem but 7. xi. 1974 - 3. xii. 1974, E-Y: 452 (TMSA). 2 ♀ idem but 4. xii. 1974 - 1. i. 1975, E-Y: 501 (TMSA). 2 ♂, 1 ♀ idem but 2. i. 1975 - 19. i. 1975, E-Y: 534 (TMSA). 1 ♂, 3 ♀ idem but 19. i. 1975 - 7. iii. 1975, E-Y: 551 (TMSA). 3 ♂ idem but 8. iii. 1975 - 4. ix. 1975, E-Y: 727 (TMSA). Omaheke Region. 1 ♂ Gobabis, [22 ° 27 ′ S 18 ° 58 ′ E], iii. 1921, Dr Knobel (SAMC). Hardap Region. 1 ♂ Zaris Ranch, Maltahöhe, 24 ° 58 ′ S 16 ° 22 ′ E, 6. iii. 1975, FC De Moor (SAMC). Karas Region. 1 ♂ Namib, Kanaan [farm, lodge], Helmeringhausen W, 25 ° 53 ′ S 16 ° 07 ′ E, 7. v. 1977, E-Y: 1323, hand-collected, dunes, S Mothlasedi (TMSA). SOUTH AFRICA: Northern Cape Province: Kgalagadi Transfrontier Park. 1 ♂ Nossob Rest Camp, 25 ° 25 ′ S 20 ° 36 ′ E, 23. i. 1994, CH Scholtz (TMSA).	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFEE3B4ED7F2F891FB34FEAA.taxon	description	Description of the holotype. Size: Length: 11.6 mm. Width: 6.6 mm. Body shape. Convex, elongate oval, elytral margins almost parallel, elytral profile convex, broadly rounded, not strongly declivous posteriorly (with no clear transition to the declivity), attaining maximum height at middle. Colour (Figs 13 – 14): Matt black. Body setae mostly brownish yellow. Head (Figs 13 – 14): Surface sculpted, punctate (medially) to rugulose (laterally), punctures close (interspaces varies between equal to or less than puncture diameters). Clypeus obtusely triangular, feebly convex, disc slightly elevated, apically deflexed and rounded; clypeal margin entire, smooth; clypeogenal and frontoclypeal sutures distinctly raised; frontoclypeal suture complete, with a low indistinct clypeogenal tubercle present on either side, tubercles punctate. Frons raised, with paired, arched, setose frontoclypeal carinae delimiting two semicircular depressions, separated by distinct elevated longitudinal median carina (ridge); carina punctate, arched carinae on frons marked by row of erect, stout seta. Genal angles obtuse; genae shape triangular, wide, recurved, genal surface depressed, coarsely sculpted; genal margin with distinct row of long thin straight setae. Eyes large and bulbous, scarcely visible in dorsal view. Antennal scape rounded, longer than wide, surface punctate, with long light brownish yellow setae; pedicel attached apically; antennal club brown, velutinous; setae around mouthparts dense, yellow to rust-brown. Labrum shape asymmetric; surface of labrum coarsely punctate, each puncture bearing a long stout seta. Pronotum (Figs 13 – 14): Pronotal width only slightly narrower than elytra, widest at base. Lateral margins broad, explanate and curved, irregular, coarsely dentate, attenuate anteriad, with fringes of broad spatulate setae; anterior angles acute, posterior angles almost right angled. Base of pronotum distinctly marginate (bordered) and projecting backward with weakly produced posteromedian lobe; base obliquely truncate laterally behind posterior angles; basal margin with dense fringe of spatulate setae; basal edge with distinct posteromedian notch (at scutellum), edge sinuate on either side of notch. Pronotal surface coarsely sculpted with discrete round punctures, irregularly spaced; punctures close (interspaces varies between equal to or less than puncture diameters), surfaces between punctures dull with superficial fine microreticulation. Median discal area of pronotum distinctly raised, broad anteriorly, narrowing slightly towards base forming crudely shaped triangular disc stretching almost over length of pronotum, somewhat flattened anteriorly and strongly declivous on sides and behind; median pronotal ridges distinct, broad anteriorly, with slight depression in the middle, parallel to median groove, ridges interrupted at approximately two-thirds of the length from the anterior margin by a depression; median groove entire stretching almost over length of pronotum, on anterior third widened into slight rounded depression, then partially constricted at approximately two-thirds of the length from the apex, thereafter widening into slight round depression; median basal tubercles distinct, slightly laterally offset (in line with second elytral costa), not fused to discal ridges; two lateral tubercles present on either side of pronotum, these tubercles almost completely fused to form a broad lateral ridge; distinct lateral carina present below lateral tubercles; all tubercles and ridges setose. Scutellum (Fig 13): Oval, longer than wide; surface smooth; apicolaterally rounded; mediobasally depressed, depression weakly sculpted to punctate; margin smooth, tomentose. Elytra (Figs 13 – 14): Elongate, length 2.5 pronotum length, lateral margins subparallel, widest at ca. 0.6 length; profile convex, attaining maximum height approximately in the middle, declivous posteriorly; elytral sides narrow; outer lateral margin dentate over basal third, thereafter margin smooth; setae on basal third arranged in clumps matching the dentation, thereafter setae arranged more uniformly. Humeral calli prominent, bearing small tubercles with short stout setae, humeral angles obtuse, blunt. Sutural margin raised, surface smooth, with low oval to elongate tubercles, widely separated and irregularly distributed along margin, distance between tubercles roughly double or more than length of tubercles; tubercles roughly of similar size for 0.75 elytral length, thereafter decreasing in size until barely discernible; tubercles smooth, dull shiny, each with a posterior setal tuft. Elytral costae distinct, even numbered costae (2, 4, 6, 8) more prominent than odd-numbered (1, 3, 5, 7, 9). Even-numbered costae with raised oval to elongate tubercles, these tubercles smooth, dull shiny, each with a posterior setal tuft; tubercles on costae 2, 4, 6 and 8 fairly evenly spaced along elytral length, distance between tubercles roughly equal to or slightly greater than length of tubercles; costae 2 and 4 with 10 tubercles for 0.75 elytral length (up to apical callosity); weak apical callosity present on fourth costal interval at top of elytral declivity. Odd-numbered costae (1, 3, 5, 7, 9) marked by small round to oval, dull shiny tubercles, each with one or two short setae; tubercles on costae 1, 3, 5 of roughly equal size and height for 0.6 elytral length, thereafter decreasing in size until barely discernible; tubercles on costa 9 much smaller, barely discernible. Costae 1 – 8 separated from intercostae as slightly elevated bands; costa 9 barely discernible as band. Intercostal surfaces strongly undulate, marked with large round to ovoid punctures (depressions), punctures separated by transverse ridges, punctures closely spaced along elytral length, distance between ovoid puncture roughly equal to the diameter of puncture, punctures on intercosta 10 (adjacent to elytral margin) larger, oval in shape; margins of intercostae 1 – 6 marked by narrow longitudinal ridges (carinae); ridges on intercostae 7 – 10 vestigial, barely discernible or lacking. Metathoracic wings: Complete. Ventral aspects: Legs (Figs 13 – 15): Procoxa narrow, flattened anteriorly, surface rugose and punctate, basal section indumentose. Profemur large, widest basally, marginate on anterior surface with row of straight, forwardpointing setae; ventral surface with regularly spaced round punctures, each with a single short seta, surface dull, with superficial reticulation; dorsal surfaces smooth. Protibia dorsally keeled, terminating with small upwardsprojecting apical process and row of short stout transverse setae; protibial apex divided into two teeth; protibial spur stout, pointed anteriad, flattened laterally, apex slightly curved, slightly longer than third tarsomere; tarsal segments smooth, tarsomeres 1 – 4 subequal in length, fifth tarsomere longest, with two simple and equal claws; tarsomeres 1 – 2 bare, without setae, tarsomeres 3 – 4 laterally with a single short seta on either side, ventrally each with a single long seta; outer tibial margin with two distinct median teeth and four to five smaller basal teeth; outer tibial surface rugose and punctate, each puncture with short straight seta; in lateral aspect the dorsal longitudinal ridge (keel) on protibia dentate, with row of long, straight setae; both the inner and ventral tibial surfaces smooth and shiny, longitudinally with two crude, shallow, almost continuous parallel grooves (formed from rows of coalesced punctures); grooves on inner tibial surface with numerous long thin setae, setae on ventral surface shorter and more sparse. Meso- and metafemora slender; anterior margins rounded, posterior margins somewhat flattened, ventral face rugose, with regularly spaced round punctures, each with single stout seta; dorsal surfaces smooth. Mesotibia and metatibia slender, outer margins dentate and with short, stout yellow setae; inner surface smooth, glabrous, upper and lower margins with long straight setae; outer margin on apex of both meso- and metatibia with row of closely spaced stout backward pointing setae (comb); inner margin of meso- and metatibia with two apical spurs, upper spur slightly longer than second tarsomere, lower spur slightly longer than first tarsomere; tarsomeres of both meso- and metatibia smooth; tarsomeres 1 – 4 subequal in length, anterior margin of each segment with four spinelike setae, two ventrally, one dorsally and one on outer margin; fifth tarsomere as long as tarsomeres 2 – 4 combined, with two simple and equal claws. Proepisterna: Surface glabrous, with fine, scattered setae, basal margin distinctly raised, marginate, margin punctate with long thin setae; disc with short medial longitudinal carina. Epipleura: Broad basally, narrowing apically, weakly concave along entire length, surface mostly smooth, mid-longitudinal row of short setae along entire length; outer margin denticulate, inner margin distinctly marginate, margin tomentose with scattered short setae. Mesosternum: Sunken below metasternite, with distinct basolateral depression on either side (visible on cleaned specimens); surface rugose and coarsely punctate on basal half (approximately areolate), each puncture with single seta; surface of distal half smooth, sparsely punctate; mesosternal apophysis prominent between mesocoxae, extending almost the length of mesocoxa, narrow (thinly triangular), pointed, with medial longitudinal ridge (keeled), surface tomentose. Mesepisternum: surface weakly undulating, rugulose, sparsely setose, distal margin raised, thick. Metepisternum: surface weakly concave along entire length, strongly rugulose, with scattered thin (shorter) and thick (longer) setae along entire length, outer margin marginate, slightly raised and tomentose. Metasternum: Convex, about 1.7 times as wide as long; surface rugose and coarsely punctate, with evenly scattered short thick setae, faintly tomentose on discal area; lateral and anterior margins marginate, raised, tomentose; distal margin smooth; metaventral plate sunken along mesocoxal margin forming a narrow basin-like depression, metasternal apophysis prominent between mesocoxae, raised, keeled-like; metaventral disc rhomboidal (discrimen), visibly depressed medially, outer margin with small median pointed triangular projection; plate strongly declivous posteriorly, transverse margin strongly sinuate on either side of median depression, declivity rugulose and without setae. Abdominal ventrites: Five visible sternites (IV – VIII); sternite IV narrow, perpendicular to other sternites, concave, surface undulate, rugose, basal margin marginate, with short triangular apophysis between metacoxae; sternites V – VII with surfaces coarsely sculpted with arch-shaped grooves, grooves forming almost concentric semi-circular pattern, surface between grooves dull shiny, remainder of surface smooth, glabrous, with row of fine punctures running parallel to the distal margin, punctures irregularly spaced, each with short seta, lateral margins setose; sternites V – VII with small lateral depressions; distal margin of sternite V straight, subparallel with distal margin of sternite IV, distal margins of sternite V and VII slightly concave; sternite VIII with distal margin convex, setose, surface finely sculpted, feebly punctate. Male genitalia (Figs 16, 20, 23): Symmetrical, slender, strongly sclerotised. Phallobase subovate in dorsal view, arched in lateral view, slightly longer than parameres. Parameres slender, long, extending beyond phallus, in dorsal aspect curved inward; parameres thin in basal half, more thickened in apical half (Figs 16, 20); apex narrowed, pointed, always projecting forward (never inward or down) (Figs 20 h, 23 s); parameres in lateral aspect slender, almost straight (never curved or convex), sides subparallel (never expanded medially; Figs 16, 23 t), apically slender, narrowed, apices short, only feebly decurved (almost straight, never strongly decurved or projecting downward as in Phoberus sulcatus), projecting forward (Fig 23 s). Phallus divided into two symmetrical lobes (Figs 16, 20); lobes broad basally, surface there slightly depressed (Fig. 20 l), lobes strongly narrowing in apical third, outside margin distinctly notched at apical third (Fig. 20 j), apex of phallus narrow, rounded, projecting upward (cf. keeled) (Fig. 20 i); inner margins of lobes feebly concave to almost straight (never strongly concave), parallel and close (Fig. 20 k); two bladelike projections and hooks visible between parameres and phallus. Variation. Size: length 10.1 – 12.8 mm, width 5.4 – 6.6 mm. Arrangement, number and shape of tubercles on the even- and odd-numbered costae variable between individuals. Secondary characteristics are limited to the protibia (if not worn down): in males, the apex of the protibia is bidentate (teeth narrow, equal); protibia of males narrower than in females; females have the protibal apex not distinctly bidentate (teeth partly fused, outer tooth wide, round), protibia slightly broader than in males. No other differences were observed in the external morphology of either males or females.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFEE3B4ED7F2F891FB34FEAA.taxon	diagnosis	Diagnosis. Phoberus erwinhaafi Strümpher, sp. nov., is most similar to P. instabilis revalidated and new combination, but males of the new species can be distinguished from the latter by the following combination of characteristics: the parameres in lateral profile are more slender, fairly straight and not expanded medially (Fig. 23 t); apices are also shorter and straighter (Fig. 23 s); Furthermore, in Phoberus erwinhaafi Strümpher, sp. nov. the lobes on the phallus are broad and the inner margin weakly concave and close, not straight as in P. instabilis revalidated and new combination (compare Figs 19 g and 20 k). Also, see Table 1 for additional characteristics.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFEE3B4ED7F2F891FB34FEAA.taxon	distribution	Geographic distribution. The species is known from Namibia and the Kgalagadi Transfrontier Park in the Northern Cape province of South Africa (Fig. 24).	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFEE3B4ED7F2F891FB34FEAA.taxon	biology_ecology	Natural history. There are collection records from carrion (meat). Based on label data, Phoberus erwinhaafi Strümpher, sp. nov. is primarily active in the austral summer (September to May). Immature stages are unknown.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
039B87A2FFEE3B4ED7F2F891FB34FEAA.taxon	etymology	Etymology. This new species is named after Dr Erwin Haaf, German entomologist associated with the Museum G. Frey (Germany), in recognition of his significant contributions to the study of Trogidae.	en	Strümpher, Werner P., Huchet, Jean-Bernard, Costa-Silva, Vinícius Da (2025): Phoberus sulcatus (Thunberg, 1787) sensu lato: in-depth morphological analysis of male genitalia reveals three cryptic species (Coleoptera: Scarabaeoidea: Trogidae). Zootaxa 5660 (3): 301-329, DOI: 10.11646/zootaxa.5660.3.1, URL: https://doi.org/10.11646/zootaxa.5660.3.1
